Abstract: Farming was first introduced to southeastern Europe in the mid-7th millennium BCE - brought by migrants from Anatolia who settled in the region before spreading throughout Europe. However, the dynamics of the interaction between the first farmers and the indigenous hunter-gatherers remain poorly understood because of the near absence of ancient DNA from the region. We report new genome-wide ancient DNA data from 204 individuals-65 Paleolithic and Mesolithic, 93 Neolithic, and 46 Copper, Bronze and Iron Age-who lived in southeastern Europe and surrounding regions between about 12,000 and 500 BCE. We document that the hunter-gatherer populations of southeastern Europe, the Baltic, and the North Pontic Steppe were distinctive from those of western Europe, with a West-East cline of ancestry. We show that the people who brought farming to Europe were not part of a single population, as early farmers from southern Greece are not descended from the Neolithic population of northwestern Anatolia that was ancestral to all other European farmers. The ancestors of the first farmers of northern and western Europe passed through southeastern Europe with limited admixture with local hunter-gatherers, but we show that some groups that remained in the region mixed extensively with local hunter-gatherers, with relatively sex-balanced admixture compared to the male-biased hunter-gatherer admixture that we show prevailed later in the North and West. After the spread of farming, southeastern Europe continued to be a nexus between East and West, with intermittent steppe ancestry, including in individuals from the Varna I cemetery and associated with the Cucuteni-Trypillian archaeological complex, up to 2,000 years before the Steppe migration that replaced much of northern Europe's population.

Yes, looks amazing this one too. I just don't know how they manage to publish two huge papers on the very same day (after 2 years waiting). We'll get crazy trying to get our heads around 400 new samples and following comments about both papers.

No, I'm not complaining. This is a feast. But it'll take time to digest all this new data. I'd recommend not to shoot too fast on the comments. With all this data, better to take is easy. At least it's what I'll try to do.

Just want to say thank you for the link! Since receiving my mtDNA results, I've been driven to research my ancient Grandmothers migrations & Cultures! The hill I live on has 3 chert "ovens" and a chert quarry( and it's crumby chert, has to be heat treated in order to make points) The Native Americans that mined this flint were called Archaic Indians. I started thinking about where my ancient folks lived and knew it wasn't here! Now, I know that my ancient relatives were distantly related to these Native Americans (A.N.E.) thousands of years ago in Siberia!

If I identify them well in supplementary table I then these are all pretty early samples (by the time span of Comb Ware), pretty much contemporary to Samara_Eneolithic or even earlier. Samara_Eneolithic is also overwhelmingly EHG. There is special isolation here, just a fresh westward population expansion from the NE European forest zone.

455 An alternative hypothesis is that the ultimate homeland of Proto-Indo European languages456 was in the Caucasus or in Iran. In this scenario, westward movement contributed to the457 dispersal of Anatolian languages, and northward movement and mixture with EHG was458 responsible for the formation of the population associated with the Yamnaya complex. These459 steppe pastoralists plausibly spoke a “Late Proto-Indo European” language that is ancestral tomany of the non-Anatolian branches of the Indo-European language family.52 460 On the other461 hand, our data could still be consistent with the Steppe-Balkans-Anatolia route hypothesis462 model, albeit with constraints. It remains possible that populations dating to around 1600463 BCE in the regions where the Indo-European Luwian, Hittite and Palaic languages were464 spoken did have European hunter-gatherer ancestry. However, our results would require that465 such ancestry was not ubiquitous in Bronze Age Anatolia, and was perhaps tightly linked to466 Indo-European speaking groups. We predict that additional insight about the genetic origins467 of the potential speakers of early Indo-European languages will be obtained when ancient468 DNA data become available from additional sites in this key period in Anatolia and the469 Caucasus

Looks like this is the source of that Krause Map. They think CHG is PIE.

We predict that additional insight about the genetic origins of the potential speakers of early Indo-European languages will be obtained when ancient DNA data become available from additional sites in this key period in Anatolia and the Caucasus

But I'm pissed off that they mentioned Iran in that part, because there's no ancestry from ancient Iran on the Bronze Age steppe.

It doesn't seem that the results are against a Steppe PIE Homeland, quite the contrary. But, without some samples from Anatolia in a clear IE Anatolian context, it will be difficult to be sure. But if R1 a/b is rare in the Caucasus, as it is likely, the hypothesis of CHG = PIE would become very difficult to defend. It is hard to imagine a very patriarchal/patrilocal/patrilienal society, as deduced by linguists for PIE, tansmitted by women, with a very long duration (at least hundred of years, as CHG arised from 30% in Khvalynsk to 50% in Yamna). Not very convincing.

It is very likely that some of the authors do actually think that CHG is a possible source for PIE, or else they would not have included that part, since this hasn't even been through the peer review process yet.

I think that idea is weak, and there hasn't been any DNA from the Anatolians who actually wrote things in IE.

Clearly, language is not 100% tied to genetics, so the early IE situations may just be unlikely or unusual cases.

Well it fails the Balkan stage It fails the Anatolia stage, or is best inconclusive South Asia will certainly be key evidence But at present it only seems that NW indoeuropean expanded from the steppeAt least you have that mate. :)

Well, this paper just adds the the ?? in my personal research on mtDNA H6a1a. H6a1a is a PC Steppe lineage to CWC. My 700 AD matches are mostly in the Balkans. So it looks like Yamnaya/Samara DNA was present in the Balkans during the Bronze Age.....

In eastern Europe we479 document the appearance of CHG/Iranian Neolithic ancestry north of the Black Sea, and its480 eventual extension as far north as the Baltic. In some ways, this expansion parallels the481 expansion of Anatolian farmer ancestry into western Europe although it is less dramatic, and482 several thousand years later.

Moreover, whileBronze Age Anatolian individuals27 448 have CHG / Iran Neolithic related ancestry, they haveneither the EHG ancestry characteristic of all steppe populations sampled to date20 449 , nor the450 WHG ancestry that is ubiquitous in southeastern Europe in the Neolithic (Figure 1A,451 Supplementary Data Table 2, Supplementary Information section 1). This pattern is consistentwith that seen in northwestern Anatolia11 and later in Copper Age Anatolia23 452 , suggesting453 continuing migration into Anatolia from the East rather than from Europe.

To be neutral you can't connect autosome with Hgs directly . But obviously we are not worried about M417- clines , they were there all over Eurasia from Meso to Early Neo , we will have to see how the pattern of M417 + emerges.

I also think Meso India will also show lots of R1a M417 - , but again it will not matter much to the IE question, to which M417 can be somewhat faithfully connected.

Now it turns for the Mycenaean and Maykop , very very crucial and of course India.

As I already said, I don't care whereas it's CHG or not. I have no specific commitment on the subject.

But clearly, it is not very parsimonious.

Now, we need to find a solution for the Anatolian branch of IE. The problem is that we don't have samples tested, and it could be very difficult to have some. As you know, fire is not very friendly to DNA. And Barcin seem to be connected to a larger cultural complex in connection to previous Late Neolithic culture and Thrace.

In fact, if we could obtain some later samples, from the Hittites period, it would probably settle the thing. If the Hittites themselves were not too much admixed (but I have a bad feeling on this). But the burial practice is a central difficulty here.

By the way, a route to the Balkans is not the only solution, and this question is secundary to the question about Hittites ethnogenesis. It will be time to try to find the route if they have some Steppe admixture.

Also the Yamnaya type ancetry that S Asiana show is a ''Pseudo-Steppe ancestry'' , created from pre existing local genome . This will be crystal clear from aDNA of India . Of course we need Neo to Iron coverage .

Earlier you found that Lengyel_LN is the best reference population for the source of "farmer" ancestry in later mixed populations. It would be interesting to redo those tests with GAC as a possible source.

We are likely facing a huge survivorship-bias when it comes to the deep roots of language families. Probably neither IE nor Uralic is old enough to be the language of the EHG as it is. Comb Ware and Yamnaya separated from the post Ice Age re-population of East Europe by more time than they separated from us. There is no good reason to assume that there was no linguistic diversity in such a huge region after such long time (even assuming that the re-population had only one relatively homogeneous source, and that is already a strong assumption). We do not even know the geographical extent of the "EHG" into the east actually.

Because the early Indo-Europeans were highly patriarchal and patrilineal, with a vocabulary to match.

And what makes CHG people less patriarchal?? , the indication from autosome is that CHG works for PIE , while ANE related ancestry both in the Subcontinent and E Europe can be an archaic hunter-gatherer artifact not related to PIE at all , but obviously it is just simple thinking, in truth we don't have a clue of what languages who spoke...

I know you are reading comments at least the first few , you guys should account, that Anatolian branch is quite older given their language , so they should be there from before 2000 BC , well before actually.

@Romulus, it would seem Varna had some steppe ancestry according to the new paper, although it may have been spotty and not homogeneous throughout the population.

"In a few individuals from southeastern Europe, we find evidenceof steppe-related ancestry far earlier (defined here as a mixture of EHG and CHG similar to the genetic signature of individuals of the later Yamnaya; Figure 1B,D). One individual (ANI163) from the Varna I cemetery dates to 4711-4550 BCE, one (I2181) from nearby Smyadovo dates to 4550-4450 BCE, and a third individual (I1927) from Verteba cave, associated with the Cucuteni-Trypillian complex, dates to 3619-2936 BCE. These findings push back by almost 2000 years the first evidence of steppe ancestry this far West in Europe, demonstrating the resumption of genetic contact between southeastern Europe and the Steppe that also occurred in the Mesolithic."

Lines 445 to 460 need to be deleted. They seem to be disconnected to all the evidence presented and referenced and more amazingly the fact is admitted. The author of these lines needs to give the reasons as it makes no sense in terms of IE.

this kind of diversity of R1 lineages is only known on the Iranian Plateau, East Anatolia and South_Central Asia. However only with modern samples non ancient unfortunately. More samples would be good.

Re: this paper, hmmmm.... much of the cool stuff has already been quoted (e.g. Bronze Age Anatolia), a few bits that jumped out to me that don't seem to have been commented yet:

Also: We find that four southern Greek (Peloponnese) Neolithic individuals – three from Diros Cave and one from Franchthi Cave, plus one previously published individual from Diros27 374 – are not consistent with descending from the same source population as other European farmers.

In PCA these individuals are outliers; shifted away from northwestern Anatolian and European Early Neolithic individuals, in a direction opposite from WHG. D-statistics (Supplementary Data Table 2) show that in fact, these “Peloponnese Neolithic” individuals have less WHG-related ancestry than Anatolia Neolithic ones, and that they form an outgroup relative to Anatolian and Balkans Neolithic populations, suggesting an independent migration into Europe from a population that split off from the ancestors of the northwest Anatolian individuals from which we have data.

They touch on various models for this, including Levant Farmer->Peloponnese, but, one conclusion they don't touch on would be WHG ancestry moving into Turkey, with less admixed "East Mediterranean HG" (EMHG, I'll coin it) preserved in the Peloponnese.

As a mirror to the above, Iron Gates as KO1 like? (majority WHG, minority EHG + AN) "Modeling Iron Gates hunter-gatherers as a mixture of WHG and EHG (Supplementary Table 3) shows that they are – as expected given their geographic location and the hunter-gatherer ancestry cline – intermediate between WHG (87%) and EHG (13%). However, this qpAdm model does not fit well (p=0.0003, Supplementary table 3) and we note that the Iron Gates hunter-gatherers carry mitochondrial haplogroups K1 (8/36) as well as other subclades of haplogroup U (27/36) and haplogroup H (1/36).

This contrasts with WHG, EHG and Scandinavian hunter-gatherers who almost all carry haplogroup U5 or U2. Therefore the Iron Gates hunter-gatherers have ancestry that is not present in WHG or EHG. This suggests either genetic contact between the ancestors of the Iron Gates population and hunter-gatherers from Anatolia, or that the Iron Gates population is related to the source population from which the WHG split off during a post-LGM re-expansion into Europe. "

Sophisticated late Mesolithic Lepenski Vir culture? Anatolian Farmers - In contrast, the two individuals (I4665 and I4666, dated to 6205-5907 calBCE and 6222-5912 calBCE respectively) that we sampled from Iron Gates site of Lepenski Vir are genetically farmers rather than hunter-gatherers, despite having been buried in the local Mesolithic tradition.

Also, interesting to see some "Steppe" ancestry in the Balkans much earlier than in Hungary...

Not so sure about the sex-biased admixture results. Nick Patterson has broached concern about our methods on these before... But it would at least fit with the greater surgence of I2 in MN populations.

The three Bronze Age Anatolians are from the district of Atabey in the province of Isparta. In later times the Anatolian IE language Sidetic was spoken there, but this early it's quite possible that the population was still Hattic or something related.

The authors dismiss the Balkan route for Anatolian languages out of the reasont that even during the Bronze Age Steppe ancestry was very rare and appeared only sporadicly across the Balkans with no connection to a culture. So like a few individuals got "lost" there. However despite this R1 linages all the way to Mesolithic/Paleolithic times. The Balkans are rivaling Mal'ta.

Pigmentation: Iron Gates HG were more or less WHG type (80:20) to the extent they find that's a valid model. Derived SLC24A5 hits around 0.6 in this population (higher than SHG). Derived SLC45A2 hits around 0.2 as well, in contrast to 0.05 in WHG.

Adds some confusion about why the WHG did not have this variant.Likewise, the EHG linked populations seemed to have derived SLC45A2 at high frequencies. Not Ukraine HG (40:60 WHG:EHG) with aroun 0.2.

(Adds to a muddle to me about what the true pigmentation of Euro HGs was, and whether we really can impute a dark skinned phenotype to Western European WHG with confidence. We should be able to, as we have models of the trait in animals and admixed recent humans. OTOH, we are talking about a population which contributed perhaps 10% of ancestry at most to present day Europeans, and even then possibly only in the West. so maybe we're missing some variant that is important to them, but contributes less to present day variation).

On another note: KITLG SNP rs12821256 that is associated with – and likely causal for – blond hair in Europeans4,5 is present in one hunter-gatherer from each of Samara, Motala and Ukraine (I0124, I0014 and I1763), as well as several later individuals with Steppe ancestry. Since the allele is found in populations with EHG but not WHG ancestry, it suggests that its origin is in the Ancient North Eurasian (ANE) population. Consistent with this, we observe that earliest known individual with the derived allele is the ANE individual AfontovaGora 3

Lactase: The WHG individual Ibousierres-25 appears to carry the derived allele at the SNP rs4988235 that is strongly associated with lactase persistence in present-day Northern Europeans. Seems a bit WTF, esp. given paper (last year?) imputing this to an EEF background, and as no dairying in the Mesolithic, but if it somehow gave another advantage, and then was selected later for its lactase benefit.

I cant believe these guys didnt sample the Mycenaeans or Minoans. What the hell? Will we ever get them? On another note..The Neolithic Peloponnese looks pretty similar to The Neolithic farmers from Greek Macedonia..only more Southern shifted/slightly less WHG shifted.

Very interesting that the Bronze Age Anatolian from near Isparta has Y-J1a. As mentioned, these samples could be speaking a substrate language to Luwian and later Lycian. There is much J1(xP58) in Western Anatolia in modern populations.

Well Simon, what can I say? Data seems to indicate that the extra CHG and affinity to modern Middle Eastern populations in the Balkans today can't be explained by dynamics taking place prior to the Late Bronze/Iron Age. This reinforces the possibility of migrations ensuing from the Bronze Age collapse in the East Med as the culprit.

Btw, I am quite bummed out by the lack of samples from Southern Italy(the paleolithic one doesn't count lol) and western Balkan mountain chain(save for Govrlevo).

@Arza“Ukraine Eneolithic 3500 BCE I4110 from Dereivka looks like a modern Balto-Slav (Figure 1.)”Ukraine Eneolithic 3500 BCE I4110 from Dereivka looks very interesting. It is a pity we don’t have Y-DNA. It is much later than other Dereivka samples who look like a bunch of HGs.

Notice that it is on the line extending from Trypillia to Trypillia outlier:

http://s22.postimg.org/inmma7oj5/screenshot_180.png

Maybe this line will lead us to some R1a PIE population from the steppe?

"And some Comb Ware were close to 100% EHG. Very interesting that they were so isolated for so long."

Isolated? From whom or what?

The gothic Pit-ware and the finish Comb-ware were quite closely related already during NME/ENE - as they both related to the Atlantic Pontus at the inning of the Finnish Gulf to the great Volga - creating the major track of travel and trade bewteen Europe and Asia. This was populated and ran by the finns and used by the goths, too - creating the Pit-Com-Ware in the middle of the ancient north.

Which may explain why the first northern farmers could move into the Bay of Riga and the Gulf of Finland - carrying hgs like R1a and U4 - migrating from Sweden and Gotland to Narva and Oleni Ostrov and the lower Volga, Srubnaya and Sinthasta. Making various styles of corded ware as well as pitted and combed ware. Just like their somewhat older Scandiavian and East-European brotherlines.

Thus there's a clsoe relation between the Scania-Motala-Ajvide population and the "EHG" (I2/I1 - U5b). Reflecting the close connection to the "WHG-motive" made from Motala-Loshbour. Which is why you find "unacounted substructures" in the rundowns using WHG and EHG both.

A similar close relationship is reflected in the "eastern" vs "western" farmers. As the Narva-pottery from Estland, with R1a1a/U4 is reflected in the CWC it's also reflected in the female part of the Yamna-horizon, otherwise carried by an eastern branch of R1b. Due to the close conection between the R1a and R1b dynasties - that just like todays farmers from Sweden (R1a) and Denmark (R1b) - kept marrying their laborous daugthers to eachothers sons.

Thus we may explain the close mitocondrial connections between the early agriculturalists of Narva/R1a and Yamna/R1b. As well as the close y-connections between the early R1a-farmers of Sweden/Norway, Baltic/Carelia an the Srubnaya, Androvno and Sinthasta.

Thus the heartland of the 8.000 year old Comb-ware-culture were at the very helm of the old highways across Eurasia, untill Mercators time marking the border between 'Europe' and 'Asia'.

"Looks like this is the source of that Krause Map. They think CHG is PIE."

Krause is completely unaware of the largest waterway in Eurasia and the Caspian-Carelian connection. Thus he is missing half the map - in order to understand anything about east-west movements - of everything - during mesolithic, neolithic, eneolithic, bronze-age and iron-age Eurasia.

In Davidskis post "Dead-cat-bounce" that became no less than embarrasing. All the more so as this connection is explaining most of his enigmas.

@Simon,"Ha, Jesus, no R1b/R1a in GAC, they were pure I2, and neither were they steppe admixed."

That's a bit curious because it theoretically could mean that the segment of Neolithic BBC/CWC ancestry that can be modeled as GAC/TRB could have happened much further east instead of North-Central Europe as is currently assumed.IOW, the CWC turnover in Northern Europe could have been much more extreme.

I think you missed the important details: steppe ancestry shows a Nadir during the early Bronze Age, just when the Kurgan invasions were meant to be happening into the Balkans. There is one middle to late Bronze Age steppe-peak in an R1a-Z93 guy, but that doesn't say anything. Prior to the Bronze Age, we see are sporadic individuals during the Chalcolithic and Mesolithic. All in all there is no denonstrable "steppe migration" to the Balkans

For heaven's sake, partiarchal people shift languages like anyone else. Look at modern steppe people.

Temper your outrage or you might get indigestion, or something even worse.

Of course we can see this is their Y-chromosomes. In other words, as a rule there's no language shift without significant introgression from the carriers of the new language.

But there's no sign of any major contribution from southern men to the lineages of Eneolithic/Bronze Age steppe clans. And this is also reflected in the Y-chromosomes of modern-day Indo-European speaking Northern Europeans, isn't it?

I'm playing now with DoHA overlay... Trypillia outlier looks like Hungary BA I1504.What is even better mysterious RISE568 apparently is also on this line. RISE568-like + Ukraine_N-like (Latvia MN?) will give Baltic BA, which fused with maybe again Hungary BA will give a cline from Czechs/Slovaks/UkrWest to Latvians.

But how all of this connect to Indo-Iranians? And what about R1a CWC and possibly Yamnaya_Ukraine (totally different cline)?

And this is genetics only... now think about languages. The only expected thing here is R1a on its way to Greece.

These guys have the same Yamnaya/WHG/EEF ratios as Northern Bell Beaker and Unetice."

Tx.

Another evidence that the major river-routes were the essential way of connecting Europe - from ocean to ocean. As from the central Baltic to the Balkanic aka Black Sea. Barry Cunliffes major work "Europe Between The Oceans" keeps going strong...

"Of course we can see this is their Y-chromosomes. In other words, as a rule there's no language shift without significant introgression from the carriers of the new language."

I never liked the patriarchal language argument for genetics. I remember a few years back when you argued that because PIE was patriarchal and because early IE groups were associated with R1a, we therefore wouldnt find any r1b in PIE. Turns out that was wrong.

Many things can happen genetically in a patriarchal society. Diversity from the get go or incorperation of other men. I dont see how a patriarchal society would preclude these things. David anthony even suggested that late PIE was very much into the idea of social contracts and steppe hospitality. This stuff incorperates men.

Now, were steppe men incorperated into PIE from CHG people... I agree with you, no way. I would expect there to be significant Ydna introgression if there was such a language transfer. But I dont see how that argument depends on PIE being patriarchal.

Genetically speaking, I think the PIE people just moved around a lot with their families and helped each other out. Hence the massive expansion of ydna R1 and steppe autosomal ancestry.

@Rob"All in all there is no denonstrable "steppe migration" to the Balkans"

Yes, if you consider an early migration around 4000 BC. But the Anatolian languages are only attested after 2000 BC, not before. It is therefore likely (and consistant with History, as related in Assyrian sources) that Anatolian IE speakers were intrusive in Anatolia in the Bronze Age. Hatti were not IE, and Hittites arose to power only around 1600 BC (and very little is know about them before, if any). Divergence between Luwian and Hittite could be anterior to their migration into Anatolia. And frankly, given what we know about Anatolian languages geography arond 1200 BC, they could have come from the East, rather than by the Balkans. The big difficulty is always the same: the IE tree says that Anatolian branch is the first to diverge from the common PIE.So, I think that the only way to know will be to find some Hittite genome, if it could be identified as ethnically Hittite (not so easy, if, as I understand, they assimilated Hatti and others non-IE).

@ EastPoleHmm, but in CWC Germany we have samples from Beaker-like to almost Yamnaya-like (I1536, I1538). They are apparently blending into local farmers.This would work flawlessly in case of hypothetical CWC Russia, but they were on the "wrong" side in Germany.

"Lactase: The WHG individual Ibousierres-25 appears to carry the derived allele at the SNP rs4988235 that is strongly associated with lactase persistence in present-day Northern Europeans. Seems a bit WTF, esp. given paper (last year?) imputing this to an EEF background, and as no dairying in the Mesolithic, but if it somehow gave another advantage, and then was selected later for its lactase benefit."

Again - don't take any archaeological textbook theories as defining facts, unless you're able to understand AND check its line of evidence. The same goes for geneticians that accept such textbooks by default - and uncritically commutes others opinons...

The term "Hunter-Gather" is already known to be inacurate - as it's excluding any kind of domestication - as herding, horticulture and agriculture.

It's a decade ago that geneticians counted 35-40.000 years as the 'most-likely' age of the split between wolves and dogs.

Since then we've had a serie of dog-skulls ated from 36.000 to 22.000 years old - from Iberia to eastern Siberia.

This is immediately contradicting the old view of paleolithic man and his capabilities. In regard to domestication as well as other skills.

Ask any zoologist about 'taming' of wild animals and he/she can list more than ten arctic species that are FAR less difficult to domesticate, than greywolfs. On top you'll find wild goats, followed by horses and reindeer. If we still had wild cows they would probably meet the same level. Then you might turn to ducks, geese and woodhens, since they all gives food and feders. Two centuries ago they could even turn elks into 'work-horses' and bears into circus-artists.

https://en.wikipedia.org/wiki/Paleolithic_dog

Using the absense of evidence as evidence of absense used to be a common sin among post-modern archeologers. The fact of the matter is we don't have any clue to the first domestication of goats and sheep - for milk and wool. But we DO know that the peoples persisting the arctic winters during pleistocene Europe would be dearly appreciating both of them. In fact we have to prove that they could even survive without them...

Thus a modern genetician have to be open to such "surprises". Just count the number of archaeologers doing "sensational discoveries" over the last two decades and it should be clear that the textbooks from the 1980-ties were WAY off in terms of scientific sense. (Metodology is part of the problem, as well as honesty and plain objectivity.)

Looking into the LP-question we know that there's a certain difference between goat-milk and cows-,ilk. People intolerant to cowmilk/cream/butter(cheese can turn to goat-milk/cheese and experience NO or less problems. Horse-milk and reindeer-milk have yet another fine-chemistry, but so far no-one have made a paper out of it, regarding LP.

What remains is that the pics of goats and horses from Lascaux MAY very well be signs of 'dear' animals as well as 'impressive' ones. Thus there's no reason to be surprised or frustrated by the notion that the LP-alleles have shown up in mesolittic contexts.

It just goes to prove that there's still is MUCH to discover about our distant past. Not the least to correct all the misconceptions that have grown out of old besserwissers and macro-sceptics, unable to accept that absense of evidence NEVER must be misused as evidence of absense.

We better enjoy and apprecieate that we finally have geneticians able to do this kind of work. I wouldn't touched a blog like this unless I did. ;-)

"I never liked the patriarchal language argument for genetics. I remember a few years back when you argued that because PIE was patriarchal and because early IE groups were associated with R1a, we therefore wouldnt find any r1b in PIE. Turns out that was wrong."

It's a fact that both R1a and b was patrilineal, patrilocal and patriarchic.

You can't be one without being both others. Without any of them you won't get any linearity at all - which is why it had to exist.

Some millenia ago everybody understood that. Which is why all old laws built on "natural principles" where the first was that 'right' was defined by birth - as the first son and daugther would inherrit distinct functions -and thus positions and duties. Depending on the size of the family and their farm the number of siblings would vary. On large and central farms, as a manor or estate, there could be 20-30 sons and daugters required to meet the demands for production, communication and trade. Besides a number of socio-political duties as a part of greater whole, as in a kingdom.

To form a law of the land you need to define HOW the next king is going to be appoingted. Thus it was ONLY the FIRST son of the present king that could get the chrown. Which explains titles like chrownprince. Now from there you can build a Constitution and thus a Constututional Tradition, which by default become a "state of being". To ensure stability, development and progress one needs a Constitutional Law, engulfing ALL men in the kingdom with rights and plights. Ensuring that only "Right" would give "Might" - rather than the inverse, where one risk ending in lawlessness anytime.

We may like or dislike political systems. But we can't use myoptic likes and dislikes to analyze motives and reasons behind historical realities.

Hittites are first attested 2300 BC bud. By the time of attestation of all other Anatolian languages c. mid 2nd millenium, they were so differentiated that they require divergence for at least 2000 years. Coupled with that. The only demographic & settlement shift occurring in Anatolia is c. 4000 BC. You should be aware of these important facts if you want to enter debate, which is why there is little wriggle room. Also, the Bronze Age Anatolians were from western Anatolia. Not many Hurrians or Hatti there I'm afraid

And yes we have steppe individuals in the Balkans too. But they just seem individual migrants and traders; and EHG ancestry existed since the Mesolithic; and Balkans seems one source for the eventual steppe lines. So there is a long history of mutual exchange there. Undoubtedly the classic Kurgan hypothesis is incorrect.

I was wrong about the Trypillian culture having R1b, but I was correct about ancient SE Europeans having R1b and of my SE European/Italy or just (SE European not any new data from Italy) refugium theory that I created due to the presence of R1b in Villabruna.

They had one western Yamna from Bulgaria on the opposite side of the Yamna influence and he had a similar I as the Russian Yamna. You can see how all Corded Ware almost (almost all NOT all) have R1a thousands of miles from each other and how Bell Beakers except those in Iberia have the same type of R1b , with even the Iberian Bell Beakers having 2/7 R1b which is a lot and enough to explain a connection (unless its V88 than that would be a coincidence). The Yamna there is no doubt in my mind had little or no R1a and were dominated by R1b-L23 (Z2105/2103)(and maybe I) and had little or no R1b-L151/P310 these groups tend to have all the same lines and even though more than just one West Yamna will be needing to confirm this hypothesis, I think it is pretty much set in stone from the pattern of everything else. Perhaps the Yamna were the ancestors of Europeans in between (Autosomes) but I seriously doubt the modern R1a and R1b comes from the Yamna. I am starting to sense a Yamnn y-chromosome ancestor confirmation bias. I would love to be a direct male descendant of the Yamna and I really hope there is a new study but it isnt looking good. And BTW if any scientist (I.E David Reich) knows I am wrong please let me know. PS was the other paper the Bell Beaker Behemoth? I thought it would be larger.

@ John Smith"I was wrong about the Trypillian culture having R1b, but I was correct about ancient SE Europeans having R1b and of my SE European/Italy or just (SE European not any new data from Italy) refugium theory that I created due to the presence of R1b in Villabruna"

To think to a refugium in Italy or nearby after Villabruna (2 May 2016) was almost easy, in spite of Richard Rocca, Richard Stevens and all the others, but to theorize that ten years before has been done only from one: me.

@davidski, Collin"I never liked the patriarchal language argument for genetics"

Also formal IE linguistics does not say anything about patriarchy and language. David is confusing these with fairy tales from Gimbutas. We have a supposedly a supra patriarchal Iranic steppe culture complete with horse riders and wht not.. give rise to Sarmatians and female amazon warriors and riders.

mongols did not impose their language despite horsing about all over. turkics imposed language in turkey but turko-mongols adopted persian and indic despite very successful patriarchal dynasties in India and afghanistan.

That 1600 BC Z-93 again is a marker of Invasion , there is no Z-93 structure in these parts , it appears as what you expect for elite invasion . Unlike India and other areas where its presence show higher depth in time and divergence .

So with that, lets see how much composition they show EEF and WHG wise from more tests .

@ Aram "GioielloWhat's are Your bets that Vucedol Z2103 is positive for Z2110?"

So far Z2110* is very rare, and before this sample from Armenia, they were all in Western Europe, even though the smal's tree has also some samples from Russia, but belonging to an unique haplotype, and another from Switzerland. Unfortunately a sample (perhaps from Kuwait), which shares with me a SNP (FGC24408), and is in the smal's tree, didn't pay the fee at YFull and has been taken off (of course I'd have paid for him the 49 dollars, but YFull didn't answer my request, and very likely for privacy reason they don't do). Of course I thought that my Z2110, which seems the oldest sample of the Z2103 because tested from Full Genome, were the witness of a Western European origin (perhaps you know that I think not only that R1b1 came from the Italian refugium, but also that Z2103 in Samara did came from West, which would have had all the upstream subclades of the haplogroup), but we need other data now. Of course I don't exclude that these Z2103 in Western Europe may have come from east, also through recent migration in Medieval times. We'll see. I have to study deeply these recent haplotypes, and above all we need other data. I'd say that it seems clear that the most part of these haplotypes are dead end lines, as we know very well, and it is difficult to find the real ancestor of the survived line, but, as I said for Villabruna, that he could be also a dead line, but demonstrated that R1b1 was a clan of many people from which only one sample survived, but there were the refugium, so I say now that we know where R1b were in old times: in Europe, from the Western European hunter-gatherers, who expanded in many directions after the Younger Dryas. For saying that they came from East or the Baltic as Rod said, we need aDNA. We have in Italy Villabruna 14000 years ago. I am waiting that some older sample is found in these places, for saying that the refugium wasn't in Italy.

To my knowledge, it's Hatti which was attested from 2300BC, not Hittites. The confusion lies in the use of the name Hatti by Hittites, because they took control of the Hatti around 1700/1600 BC. But Hattians were non IE, contrary to the Hittites.

"By the time of attestation of all other Anatolian languages c. mid 2nd millenium, they were so differentiated that they require divergence for at least 2000 years."

That's the main difficulty, and I let it to linguists. But differentiation must not to have occured in Anatolia. It could have been the result of common migration of Anatolian IE speakers in Anatolia after 2000BC. We have other examples of the sort, like with Germans in the Roman Empire. So differentiation is not synomim with early arrival.

Hittites, in the Assyrians texts, are seen as an intrusive population, before they took over Hatti. So probably recent in Anatolia.

"Coupled with that. The only demographic & settlement shift occurring in Anatolia is c. 4000 BC."

No. There are hints of population shift around 2000 BC. For example, there is discontinuity between the last Chalcolithic settlement in Barcin, and the first Hittite settlement (200 years later).

"Also, the Bronze Age Anatolians were from western Anatolia. Not many Hurrians or Hatti there I'm afraid". Clear. West Anatolia is known to have been in relation with non-IE cultures, so not the best place to find IE. That's the problem, even if I don't understand your reference to Hatti, as they have nothing to do with IE.

"So there is a long history of mutual exchange there. Undoubtedly the classic Kurgan hypothesis is incorrect." We are completly agree on that.

I've always been meaning to ask you something since you're an unofficial expert for that part of the world----why is it not a popular idea that IE-zation of Anatolia took place via a steppe-cross caucasus/armenia-anatolia route? Why is the default scenario always steppe-balkans-anatolia? This may sound like a stupid question, in which case there's no better time to address it than now. :-)

(For the record, this paper---to paraphrase the authors' own words---does not preclude the Balkan scenario since we don't have actual Anatolia_BA on hand, but only Chalcolithic Anatolian samples).

This scenario via Caucasus was never fully dismissed. Mallory kept the door open for that route. Usually You will see here and there ideas that Anatolians came from/via Maykop.

The problems of crossing Caucasus is that Western Caucasus is not really easy to cross for large masses. The eastern corridor is much easier. But crossing the eastern corridor and then moving into Anatolia is somewhat counter intuitive.

Nevertheless Cimmerians moved in that way and settled in Cappadocia. Armenians moved via that route if Late PIE was in Steppe. Mitanni could have crossed it and moved to Syria/Kurdistan.

There was also proposals of Hittite moving via East Caucasus but now it is considered outdated.

Anyway we need samples not only from Hittite empire period. But also much earlier at last from Anatolia EBA to see what was happening. The idea Hittite came is really not realistic from linguistic and archaeological point of view.

Not time to look at all yet, but a few things that I didn't see too explicitly in the comments:

- The Anatolian Bronze Age samples are said to be published in Lazaridis 2017. So that must be an upcoming study with the other Asian samples (Maykop, BMAC, Swat Valley?). These samples have a strong shift towards Iranian ancestry (Supplementary Table 2), which together with what they write in their conclusions does seem to support that they're moving to a different model with a homeland in Asia. Let's see.

- Balkans Bronze Age: Yes, some variable steppe admixture, but no R1b and just one R1a outlier. In the single Iron Age sample, steppe ancestry falls to half, and WHG ancestry to 0.2% (Sup. Table 4).

- From Sup. Table 5, it seems that the autosomes vs. X chromosome estimates using qpAdm don't really work well, so we can't get good clues there about the origin of R1b. Strange that with all these samples and the ones from the BB paper the origin of R1b-L23 (and L51) is still unknown. And still no Neolithic Italian samples...

We'll need more time to look into all of this, but it's strange that so many samples didn't bring much more clear answers to many questions. Sampling looks a bit random, probably because of what Nick Patterson explained. I hope that Lazaridis 2017 paper will settle more questions.

Excuse my butting in too. I'm not sure about the relative popularity of steppe->Balkans->Anatolia vs steppe->Caucasus->Anatolia (and let's not forget Armenian Highlands/Transcaucasus->Anatolia outside a steppe scenario for 'Indo-Hittite' or even Indo-European), especially on the internet if that's what you're particularly referring to, but both scenarios have been proposed quite a bit as far as I know. I can't recommend anything really specific right now but the Encyclopedia of Indo-European Culture by Mallory and Adams summarizes most scenarios in their section on the Anatolian languages. Trevor Bryce seems to remain agnostic on the issue of their ultimate origins too while referring to discussions of all these scenaria.

As for the new Anatolian samples apparently showing no EHG, they are called "Bronze Age" but there doesn't seem to be an exact date for them, or I missed it. Based on the kinda schematic Figure 1, they seem to date in the 2700 to 2200 BCE range (so late Early Bronze Age). The coming Lazaridis et al. paper on them should clarify these things. We do need the Anatolians in Central Anatolia by ~2000 BCE at the latest though and maybe even as early as the 4th millennium.

We really need more sampling, especially to cover the post-Neolithic gap in the south Balkans and the Aegean (the complete lack of such samples in this study from that area was disappointing) and the area from western Anatolia to Armenia. A related migration problem, as I'm sure you're aware, is the one that brought (pre-)proto-Greek to Greece. We still can't tell for sure if it has Balkan or Anatolian associations, though in general the former is preferred by more people. But there seem to exist potential associations, if not migrations, between Anatolia and eastern/Aegean Greece around that period anyway even if not connected to the future Greek language but something else and it'll be nice to see which theories prevail and which fall.

The CA and BA Armenian samples are maybe an interesting comparison to Anatolia considering some (EBA) basically have no steppe ancestry while others (CA + MLBA) reach up to some 20-30+%. We obviously don't have the final picture just yet considering all this heterogeneity and migrations at the time.

Also, in reference to your comment in the other thread, E1b has been found in Sopot-Lengyel Hungary and Cardial Spain too. The new sample is the oldest but the third ancient one found so far in Europe as far as I know.

Middle and Late Bronze Age/Iron Age are the time when Steppic penetration in form of Iranic Cimmerians, Scythians etc are attested in and around Armenia. No wonder to find Steppic admixture than. What is more important for the origin of Anatolian languages is EBA/Calcholthic. And these in Anatolia as well Armenia show no sign of EHG admixture.

1) "To my knowledge, it's Hatti which was attested from 2300BC, not Hittites. The confusion lies in the use of the name Hatti by Hittites, because they took control of the Hatti around 1700/1600 BC. But Hattians were non IE, contrary to the Hittites."

Yes but by 2300 BC the land of the Hatti was the Hittites. It had already happened, the strata are quite clear at the site.

2) R: "By the time of attestation of all other Anatolian languages c. mid 2nd millenium, they were so differentiated that they require divergence for at least 2000 years."

F: "That's the main difficulty, and I let it to linguists. But differentiation must not to have occured in Anatolia. It could have been the result of common migration of Anatolian IE speakers in Anatolia after 2000BC. We have other examples of the sort, like with Germans in the Roman Empire. So differentiation is not synomim with early arrival. "

Aren't you a linguist ? :)Hold on, but that's a bit unlikely, I mean how did it turn out that every one that went out happened to speak Anatolian, but were otherwise already distinct languages. And how is it IE reached NW Europe as a common Italo-Celtic-Ligurian etc clade, one Balto-Slavic, and one or two at best I-A migrations, but 5 for Anatolia ?And didn;t the GErmanic migrations into the ROman Empire result in language loss ?

3) "Hittites, in the Assyrians texts, are seen as an intrusive population, before they took over Hatti. So probably recent in Anatolia."

Well it actually says that they took over Kanesh specifically. That they took over all of central Anatolia at the same time seems to be your interpolation.

4) "There are hints of population shift around 2000 BC. For example, there is discontinuity between the last Chalcolithic settlement in Barcin, and the first Hittite settlement (200 years later).'

I agree, there are indications of some conflicts and changes at this time also. But I think the major change was in c. 4000 BCs.

5) R: "Also, the Bronze Age Anatolians were from western Anatolia. Not many Hurrians or Hatti there I'm afraid".

"F: Clear. West Anatolia is known to have been in relation with non-IE cultures'"

Yes i think these were the maritime Aegean cultures which were non-IE, eg Minoans, etc. I think mainland Anatolia was solidly IE. A lot of the supposedly non-IE substrata in Anatolian has been overestimated. Virtually all their institutions appear to be native Luwian, Hittite, etc.

I suspect whatever is in print with Lazaridis et al. will outline West Asia more clearly, including Majkop perhaps. I think perhaps for now I should try to absorb more of what this means for Europe.

Grow up Dave, this signal you are getting, could and should very well be CHG ancestry in EHG(Basically shared ancestry). You very well know but of course it goes against your believes/wishes. The peer reviewed studies show clearly that Armenia EBA can be modeled as a mix of Anatolian_CHL, Iran_CHL with CHG like ancestry.

Are the R1a,R1b and I2a in Vasil'evka and +-2000 years later in Derievka related to each other ? Does this point to Genetic continuity within that area ? If so, could it not point towards the formation of Proto-Indo-European between 7000 and 4000 BCE within that area ?

I'm sorry because you are making a confusion between Hattians and Hittites, for obvious reasons (confusion made in the XIXth century, and because the take-over of Hatti territories by Hittites didn't change the name of the place, stil designed as the Land of Hatti). But Hattic, the langue of Hattians, is partially known even if it's an isolate (as Hurrian), and it is not a IE language (as an example, it was agglutinative, so not fusional as the Hittite/Nesite and the large majority of IE languages).

Frankly, there is no doubt about this, and very little that Anatolia was overwhelming Hattian and Hurrian in the late IIId millenium BC.

The relation of the birth of the Hittite kingdom is very interesting, as the known text show a conflict between two branchs of a same royal house, one of them with Hattian names, the other with IE names. During the conflict, Kanesh was destroyed and Hattusa became the capital of the new, unified kingdom (it's only a resume of the text itself). If you read between the lines, it seems there was a conflict between Hattians (non IE) and Hittites (IE). The Hittites won, and absorbed the Hattians (Hattic became extinct shortly after) and a large part of their culture.

So, it does seem that some IE speakers (of the Anatolian branch)became sufficiently numerous in Kanesh to impose their language to their Hattian king, and finally conquered all the land of Hatti, before conquering nearly all Anatolia, and absorbing the defeated, and imposing their culture (admixed with Hattian culture) and language.

The fondation of the Hittite Kingdom is usally dated from the XVIIth or XVIth centurirs BC.

The Kanesh IE pety kingdom could have been relatively small (probably a city-state), and doesn't need any mass migration.

A comparison could also be make with Rome(1000 years later) and its relations to Etruscans (roman royal house was Etruscan, and Romans adopted many cultural customs from them).

If you making a confusion between Hattic and Hittite/Nesite, between a non-IE language, and an IE language, how could you try to find a solution to the Anatolian languages?

Anatolian samples were probably predating the arrival of IE speakers in Anatolia. They were probably Hattians or related to them (so spoke probable Hattic a non-IE language).There is no reason to think that IE populations were already present in the Chalcolithic or EBA. There is no sign of them in archeology or in the few texts we have.

"I'm sorry because you are making a confusion between Hattians and Hittites, for obvious reasons (confusion made in the XIXth century, and because the take-over of Hatti territories by Hittites didn't change the name of the place, stil designed as the Land of Hatti).But Hattic, the langue of Hattians, is partially known even if it's an isolate (as Hurrian), and it is not a IE language (as an example, it was agglutinative, so not fusional as the Hittite/Nesite and the large majority of IE languages)."

No, I am not . The "Land of the Hatti" was continued to be used in the Hittite period. The Hittites arrived c 2300 BC because there are no subsequent destructions. This has been physically explored.

From Bryce:

""But our main focus will be on central Anatolia. Here developed anumber of prosperous settlements, presumably the nuclei of smallkingdoms, in a region extending from just below the southern bend ofthe river now known as the Kızıl Irmak (Red River)3 northwardstowards the Pontic zone along the southern shore of the Black Sea.

Prominent amongst these settlements was the site now known as AlacaHo¨yu¨k, which lies some 180 kilometres north-east of the modernTurkish capital Ankara. The settlement which was founded here in theLate Chalcolithic period reached its peak in the Early Bronze II phase, asillustrated by its thirteen ‘royal’ shaft graves and their spectacular gravegoods, generally dated to c.2300–2100. But it continued as a flourishingcommunity to the end of the Late Bronze Age. Its Bronze Agename is unknown, though Hittite Arinna, city of the Sun-Goddess, ispossibly to be identified with it. Other important settlements wereHattus, the site of the later Hittite capital Hattusa, Alis¸ar lying 80kilometres to the south-east of Hattus (and probably the ancientAnkuwa6), Zalpa, which lay in the Pontic region, and Kanesh.The last of these, Kanesh, is located in the fertile Kayseri Plain justsouth of the southern bend of the Kızıl Irmak river on the site with amound now known as Ku¨ltepe. A Chalcolithic site in origin, it has ahistory of continuous occupation down to the Roman period. But itsmost flourishing phase occurred during the Early and Middle BronzeAges. "

And "The name of a king of Kanesh called Zipani figures in a well-knowntradition which deals with a rebellion of seventeen local rulers againstthe Akkadian king Naram-Sin (c.2254–2218),"

"Contrary to earlier views, there has now developed a consensus among linguists that entry of Indo-European speakers into Asia Minor was much earlier than previously assumed. See Melchert (2003a: 23-6) with references to Carruba (1995), Oettinger (2002a) and others, and also Lehrman (2001: 116-7) and Yakubovich (2010: 6-7). The gist of the argument is that the attested degree of differentiation of the IE Anatolian languages such as Hittite and Luvian already by the beginning of the second millennium requires at a minimum that their divergence from Proto-Anatolian began by the middle of the third millennium. It may easily have begun as early as the end of the fourth."(The Position of Anatolian; Berkeley)

@RobYou do realize that your answers relie on pure hypothesis?And as I said, that Hittites adopted a large part of Hattians culture, so why should they destroy everything? Earlier settlements were not destroyed, so what? Is there a clear connection to Hittites? No? So it proves nothing.

By the way, you are answering to facts (Hattic language, texts of different origins) with mere suppositions.

There is simply no proof of presence of IE in Anatolia in the IIId millennium. Deal with it, or find something solid.

Saying simply that there is a consensus about something leads nowhere. There was a consensus about Cultural diffusion and Paleolithic continuity some years ago. What happened?

An early divergence of Anatolian languages is likely, but it doesn't need to have happened in Anatolia. It's only because Anatolian languages are found in Anatolia around 1300BC that seems logical. But it's purely conjectural.

Thanks again Davidski for the spreadsheet! (interesting, an mtDNA RO in Yamnaya) Wasn't mtDNA RO featured in a recent article on a Glacial Refugia in the Saudi Arabian Peninsula? I find the that interesting because the Saudi Arabian Peninsula mtDNA H lineages are mostly H2 & H6. And mtDNA H2 & H6 show up in Yamnaya Culture results.

That's why I mentioned "~2000 BC". Something past that rough date can't be argued considering Hittite names from Kanesh start to appear in Assyrian documents. It's not unlikely they were present before that but a direct linguistic attestation is an absolute and inarguable terminus ante quem (so it doesn't make us retread arguments that far more qualified people have argued at this early stage of post-Neolithic aDNA sampling from the area).

We really don't know that pre-Greek/Mysian/Thracian/Phrygian/whatever-you-like Anatolia (so 15th-12th centuries on) will turn out to be that 'steppe' overall, anyway.

Completely agree. I want to be clear about my position: I am not saying that no IE population existed in Anatolia in the IIId century, or that the split didn't happen there.

But, given that the story of this specific branch is quite different from the other ones, we need to keep our mind open on all possibilities.

By the way, there are some facts:- Anatolia was not fully Indo Europeanized even in the late IId millenium BC, and some non-IE were prominant in the early IId millenium BC,- No Steppe admixture has been found in Chalcolithic or EBA Anatolia,- So far, all IE population tested did have some level of Steppe Admixture,- an introgression from the East (CHG + Levant_N) seem clear in the IIId millenium BC,- the IE languages in Anatolian are only attested after a. 2000 BC.

So, from that, the rest is purely conjectural.

The only way to find out if the Anatolian branch came from the Steppe is to find samples in a clear IE context. The problem is, Hittites burned their dead, and DNA doesn't resist fire.

So, I think that we must put everything on the table, and test every hypothesis. That's all.

@JV, Most of the IE-looking uniparental markers in Saudi Arabia, Yemen, Oman and the Gulf principalities come from Persian heritage in many Shia Arabs. OTOH, Iraq, Syria, and surrounding states is going to be a totally different situation since H* likely arose there.

I think they don't really mean Basal Eurasian in a strict sense there. It's more a population that is basal to Anatolia Neolithic. It's labeled as "an1" in the graph and it donates 74% to Peloponnese Neolithic and 71% to Anatolia Neolithic. The rest in both populations is WHG (26% and 29% respectively). So overall they're quite similar in that graph, only a tiny bit more basal in the Peloponnese.

The D-stats show that the shift in Peloponnese is towards Iran Neolithic, actually.

@Folker

So far, all IE population tested did have some level of Steppe Admixture

There are really not many ancient IE populations tested to date. I can only think of the Roman age Britons, who we know spoke Celtic (but that was a rather recent shift). If you mean modern populations, then that's rather uninformative. All West Eurasians are intermixed.

the IE languages in Anatolian are only attested after a. 2000 BC

Yes, no IE language is attested before 2000 BC, anywhere. These Bronze Age samples from Anatolia are by far the closest in space and time that we have to early (attested) IE speakers. That's something. It's clearly not definitive, but it's what we have so far, and it's already insightful. No reason to dismiss it with more speculative possibilities based on non existent data.

BTW, there are 3 samples. One is older, but has very low coverage (0.28x) so not much influence in the relevant stats that show a shift towards Iran_N. It's from 2836-2472 calBCE. A second one has a good 1.98x coverage and it's from 2558-2295 calBCE. The third one has the highest coverage at 3.69x and it's just dated as 2500-1800 BCE.

This paper hit home because im mostly british isle and 1) WHG might be ice age balkan. 2) verified that EEF got its WHG in meso balkans. 3) confirmed the British were replaced by BB, eliminating virtually all local WHG. 4) oddly enough, some extant WHG may be from Eastern Europe.

Hell, the same is even true for Poland. Most Polish WHG is from Balkans (via EEF) and Easter Europe via (CW). EHG is from East of Poland, while CHG and Northwest Anatolian are from the edges of East Europe.

This is all reinforced by Ydna. The Mesolithic Baltic, Balkans, and Steppe are all loaded with R1B. Same for R1a just shifted a bit to the east. Really strange when you think about it.

"* Early European Farmers are mostly mix of Mesolithic Balkan and Neolithic Northwest Anatolia."

Where did you got that from? The papers say nothing like that.

Early European Farmers are 95% Anatolian Neolithic all the way from Central Anatolia. Thats what the data shows.

What the new papers say is that Anatolian_Farmers merged with local WHG groups on the Balkans and that there were several waves of farmers.

The paper states that WHG might be from a Balkan refugium but the WHG like ancestry in Levant Neolithic and Anatolian Neolithic is not necessary from the Balkan but stems from a common ancestor with WHG.

There was R1b Z2103 in BA Balkans. Specifically in Vucedol culture. 50% from two male samples. If You add 3 previously known cases of male Vucedol You get 5 males with 60% of R1b .Their autosomes live little doubt that they are from Steppe.

The origin of both L51 and Z2103 in Europe is solved. Both they entered from Steppe. Thus L23 level formed in Steppe.

What is that IJ doing at Volniensky, Vilnianka ? Was it not a good sample ? If it was actually J could it explain from where CHG entered the Steppe ? Did some I2a in Southern Ukraine carry some CHG ancestry ?

Apparently Yuri Rassamakin suggested that Sredny Stog should be seen as four different Cultures within that area.(R1a,R1b,I2a and later G2a ?) Could this at least partly explain the Uralic, and Kartvelian connection with Indo-European ?

I wonder what this all means for R1a.. I2 and R1b seem to have long been existing in a line from the Baltic to the Balkans since the LUP, to-ing and fro-ing through the epochs. R1a appears in Karelia Mesolithic, then Ukraine "Neolithic". It is also in a couple of the Meso-Neolithic samples from around Baikal... Perhaps an Aral-Caspian centre of dispersion?

@ Aram "Alberto There was R1b Z2103 in BA Balkans. Specifically in Vucedol culture. 50% from two male samples. If You add 3 previously known cases of male Vucedol You get 5 males with 60% of R1b . Their autosomes live little doubt that they are from Steppe. The origin of both L51 and Z2103 in Europe is solved. Both they entered from Steppe. Thus L23 level formed in Steppe".

Aram, this is the way of reasoning of the two Richards without lion and heart. You are smarter. Unfortunately R-L51 and subclades hasn't been found in Samara neither in all the hundreds of samples tested Eastward Germany, and R1 (both a and b) was the Y of the European hunter-gatherers, and the Y line was above all masculine also in Bell Beakers of Germany and low Rhine: "On the other hand, hunter-gatherer Y chromosomes (haplogroups I2, R1 and C1) are much more common: 6/7 in the Iberian Neolithic/Copper Age and 7/8 in Middle-Late Neolithic central Europe (Central_MN and Globular_Amphora). Under a single pulse model of admixture, the autosomal/X chromosome ancestry proportions imply that in the central European Middle Neolithic population that shows the strongest evidence of sex bias, 35-50% of the male ancestors were hunter-gatherers, compared to 0-5% of the female ancestors" (Mathieson, lines 413-418). Hg. R1b was from Northern Italy to the Balkans to Baltic from at least 14000 years ago, and certainly R-L51 didn't come from Samara. Let's wait that Tyrrhnian Italy and the line Rhone-Rhine are tested at least as the places tested so far. Certainly R1b wasn't in Middle East, Anatolia, Caucasus.

@ Aram"Nevertheless imho it is safe to say that most of R1b Z2103 present in Balkanes are initially from Vucedol culture. And they entered via Hungarian corridor".

Perhaps the R-L23-Z2110 of Nochev, but the R-L23-CTS9219 are all recent and come very likely from West. See the said "Balkan cluster" (name mine), which is older in Italy and western Europe. The R-L23-CTS7763, formed in Asia after a migration from Samara, came very likely recently with Turks.

From a purely historical perspective, Hattusa was conquered by Hittites even later than Nesa (Kanesh). When Hittites moved their capital from Nesa to Hattusa sometime during the 16th century BC the Hattusa region was still largely Hattic-speaking as evidenced from the abundance of Hattian names and the preservation of Hattian writings from that region in the Hittite records and the ongoing conquest of that region by Hittites even after the city of Hattusa became their capital. Even the name of the city of Hattusa comes from Hattians, natives of the Hattusa region, who were clearly a non-IE-speaking people. Hittites always preserved the memory of coming from Kussara and conquering Nesa and Hattusa later, they did not regard themselves as natives of Nesa or Hattusa.

Where was the Main territory of the Vucedol Culture ? Was it in Hungary or rather more concentrated in Serbia and the Adriatic ? Wouldn't that point to R1b in Hungary being somewhat on the fringe of Vucedol and not part of the main action ?

We see with the Bell Beakers that Steppe R1b people were good adoptors and adaptors. (Maybe even Hittites ?) So wouldn't this make añ earlier R1b migration from the Steppe a bit more difficult to trace Culturally if they soaked up Cultural elements from other Cultures like a sponge ?

What is Western Yamnaya ? Isn't Western Yamnaya basically the earlier Sredny Stog ? Or were Sredny Stog basically totally displaced by Yamnaya ? If this was the case then where did those people go ?

Sredny Stog in their later phases show some elements similar to Later Corded Ware but what about the earlier phases and its probable connection to the people that lived in that area from +-8000 BCE. ? Sredny Stog came to an end at +-3500 bC. The Corded Ware Culture only emerged much later. So is Corded Ware descendants from Sredny Stog, Khvalynsk or Yamnaya ?